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37 of 40 people found the following review helpful
Format: HardcoverVerified Purchase
I read a lot, but I rarely suggest books to people I am acquainted with (you know, people get sick of that sort of thing); however, since I finished reading this book, I can honestly say that this is the one volume I have actually recommended to my friends and family. This book covers a crucial aspect of our modern life and is far-and-away one of the most indispensable pieces of scientific writing I have read to date. For example, take this quote from the Preface: "Many problems that challenge us today can be traced back to a profound tension between what is good and desirable for society as a whole and what is good and desirable for an individual. That conflict can be found in global problems such as climate change, pollution, resource depletion, poverty, hunger, and overpopulation. The biggest issues of all - saving the planet and maximizing the collective lifetime of the species Homo sapiens - cannot be solved by technology alone. They require novel ways for us to work in harmony. If we are to continue to thrive, we have but one option. We now have to manage the planet as a whole. If we are to win the struggle for existence, and avoid a precipitous fall, there's no choice but to harness this extraordinary creative force. We now have to refine and to extend our ability to cooperate. We must become familiar with the science of cooperation. Now, more than ever, the world needs SuperCooperators."

One reviewer called Martin Nowak a virtuoso, this is most certainly true, and it may even be an understatement. It would seem that Dr. Nowak has his hands in nearly every discipline and knows nearly everyone who is anyone in the scientific community. Furthermore, whether he's discussing Game Theory, Evolutionary Biology, Mathematics, Multi-Level Selection, Language, the Tragedy of the Commons, Networks, or Evolutionary Graph Theory, the writing is always vigorous, entertaining, and accessible. In essence, you could probably spend countless days reading works like: Darwin's Conjecture: The Search for General Principles of Social and Economic Evolution,Predictably Irrational, Revised and Expanded Edition: The Hidden Forces That Shape Our Decisions,A User's Guide to the Crisis of Civilisation: And How to Save it,Living within Limits: Ecology, Economics, and Population Taboos,Collapse: How Societies Choose to Fail or Succeed: Revised Edition,Braintrust: What Neuroscience Tells Us about Morality,The Extended Mind: The Emergence of Language, the Human Mind, and Culture (Toronto Studies in Semiotics and Communication), or Making the Social World: The Structure of Human Civilization, (like I have done) or, you could save yourself some time and read this one book. Martin Nowak and Roger Highfield have written an absolutely incredible book. I really can't recommend this book enough. Here is just one quote, of many, which I found to be sublime: "The story of humanity is one that rests on the never-ending creative tension between the dark pursuit of selfish short-term interests and the shining example of striving toward collective long-term goals. I believe we now understand how defection in the Prisoner's Dilemma can be trumped by cooperation. And, just as [Gustav] Mahler ends on an upbeat note, so I believe the emphasis on cooperation puts a more optimistic sheen on life than the traditional take on Darwin, which condemns all life to a protracted and bloody struggle for survival and reproduction. Mutation and natural selection are not enough in themselves to understand life. You need cooperation too. Cooperation was the principle architect of 4 billion years of evolution. Cooperation built the first bacterial cells, then higher cells, then complex multicellular life and insect superorganisms. Finally cooperation constructed humanity."

The chapters are: 0) The Prisoner's Dilemma, 1) Direct Reciprocity - Tit for Tat, 2) Indirect Reciprocity - Power of Reputation, 3) Spatial Games - Chessboard of Life, 4) Group Selection - Tribal Wars, 5) Kin Selection - Nepotism, 6) Prelife, 7) Society of Cells, 8) The Lord of the Ants, 9) The Gift of Gab, 10) Public Goods, 11) Punish and Perish, 12) How Many Friends Are Too Many?, 13) Game, Set, and Match, and 14) Crescendo of Cooperation. There are a couple of books I would also encourage the interested reader to pursue after reading this book, Peter Corning's: The Fair Society: The Science of Human Nature and the Pursuit of Social Justice and Chris Martenson's: The Crash Course: The Unsustainable Future Of Our Economy, Energy, And Environment)
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55 of 62 people found the following review helpful
on March 20, 2011
Format: Hardcover
For Martin Nowak cooperation is the master architect of evolution. This man is obsessed with the idea that cooperation is an indispensable driving force of evolution at any level - mutation, selection and cooperation. Without cooperation among RNAs in the primordial soup, you and me would be still one of them. Is he crazy? Nowak has been Professor of Mathematical Biology at Oxford, the first head of the Program in Theoretical Biology at the Princeton Institute for Advanced Study, and now he is full professor of Biology and Mathematics at Harvard University in his own institute called "Nowakia". Of his numerous papers more than 50 were published in Nature or Science. Nowak is a leading evolutionary theorist of our time. Why is he crazy for cooperation?

Cooperation has always been Nowak's main subject that he studies mostly with only one technique: mathematics. "We can capture the way it (evolution) works with mathematics, distilling its essence into the form of equations." "SuperCooperators" is the grand review of his oeuvre on cooperation, a kind of textbook that reads like a bestselling novel with a wonderfully lucid and enthusiastic style, thanks to Nowak's ghost-writer and kind of co-author ("with" instead of "and") Roger Highfield, an ingenious science writer and the editor of the New Scientist magazine. A layperson could enjoy just reading this book and finally has happened to learn most about a fascinating part of biology. Imagine all textbooks were written this way! Try this appetizer from the chapter on the evolution of language: "Gossip. Banter. Chat. Let's talk. Let's organize a colloquium. Even better, let's have a party! Language allows people to work together, to exchange their ideas, their thoughts, and their dreams. In this way language is intimately linked with cooperation. For the mechanism of indirect reciprocity it needs gossip, from names to deeds and times and places, too. Indirect reciprocity is the midwife of language and of our big, powerful brain."

We learn about five ways to achieve cooperation: (1) Direct reciprocity - Tit for Tat that becomes generous but has to give way to the new champion for playing the Prisoner's Dilemma, "win stay, lose shift"; (2) Indirect reciprocity - Power of reputation. This seems to be the most important mechanism driving our sociality, language and brains; (3) Spatial games - Chessboard of life. Cooperators can prevail by forming networks and clusters; (4) Group selection - tribal wars; multilevel (group) selection works if there are many small, isolated groups; migration - egoists infecting pure altruist groups - undermines cooperation; however, "at the cellular level, there's plenty of evidence of group selection"; (5) Kin selection - Nepotism. Cooperate with close kin and defect with strangers according to Hamilton's rule. Nowak is reluctant to list kin selection as a mechanism for cooperation: "I still believe that kin selection is a valid mechanism if properly formulated."

Kin selection and inclusive fitness theory has been one of the corner stones of our understanding of the evolution of social behaviour. Hamilton's rule, b/c>1/r, was the e=mc2 of sociobiology until very recently when Nowak, Tarnita and Wilson in an Analysis article in Nature claimed that it's theoretical basis is mistaken, Hamilton's rule almost never holds and decisive empirical tests of inclusive fitness theory have never been performed. This article induced a strong reaction in the huge community of well-established researchers that made their career with studying kin selection. In SuperCooperators Nowak explains his criticism. In "The math of kin selection" he reviews the history of the concept, from Haldane's answer to whether he would risk his life to save a drowning man: "No, but I would do it for two brothers or eight cousins", to Hamilton, Price and Maynard Smith's final version. Around explaining the Price equation, the basis of current inclusive fitness theory, we learn about the biography of Bill Hamilton and George Price, and why Price's equation is the mathematical equivalent of a tautology.

In the chapter "The decline of inclusive fitness" Nowak writes: "Equations seemed to arise out of nowhere in kin selection. There were many attempts at calculations that had no precise formulation of the underlying mathematical model ... This is a recipe for disaster", and "We found that in this special world (`inclusive fitness land') where inclusive fitness theory works, the calculations yield up exactly the same prediction as standard natural selection theory. Hence inclusive fitness theory comes up with no novel predictions or insights." Nowak offers a new model for the evolution of eusociality where relatedness is a consequence rather than the cause of social behaviour: A mutant determines that daughters just stay with the nest and thus happen to stay with their mother to help raising further offspring. However, what if they happen to stay with someone unrelated? Why do parents insist on caring for their own offspring? - because they happened to be close at birth and not because they are related? As an empiricist I see many empirical results supporting the relatedness cause: e.g., the sex ratio in singly mated ants is closer to the workers' benefit than to the queen's, it's the contrary in slave maker ants where workers are in an evolutionary dead end. I expect a new "properly formulated" theory of social evolution based on relatedness will be compatible with existing evidence and include Hamilton's rule "of thumb". It will offer a plethora of new predictions enthusiastically tested by next generation students of sociality.

Nowak strongly objects also to punishment being an effective method for promoting cooperation. "But this view is mistaken in my opinion" and he dissects Fehr and Gächter's classic paper to depict any shortcomings. On this topic the winning team "Nowak & Sigmund" seem to be split: Sigmund: "Punish or Perrish", Nowak: "Punish and Perrish". Nowak, the theorist, even performed experiments to show that "Winners don't punish." Indeed, we do not know yet, how punishing ultimately pays off for the punisher. In another experiment Nowak tried to prove that positive (reward) rather than negative interactions (punishment) promote public cooperation, which turned out to be only wishful thinking. As with his attack on the basis of kin selection theory, Nowak challenges the established view on the evolution of punishment, potentially stirring up novel yet un-thought ideas.

SuperCooperators is also Nowak's autobiography. He reveals his private life - how he coming from an all-boys school met his wife - and his way of interacting with his supporters: mountain climbing with Peter Schuster, walking through Rauriser Urwald with Karl Sigmund, playing soccer with Bob May, or dining with Jeffrey Epstein, the Wall Street tycoon, who built an institute for him, at the beach of Epstein's Carebian island of paradise. Nowak has also some personal sentences about each of his successful collaborators and some pages on how "the eternal symphony" (Mahler's Song of the Earth) carried a deep resonance for him and his work.

Finally, Nowak describes the game that all people on Earth are playing - the climate game: "unless people fully realize the extent to which the planet is in peril, people will fail to do enough to save it .... I believe that climate change will force us to enter a new chapter of cooperation." In search of a deeper concept that underpins all apparently different approaches to cooperation he presents the solution: "Corina's theorem will hold for any evolutionary process on Earth, in this galaxy, as well as all the others, from those nearby to agglomerations of ancient stars that lurk in the faintest, farthest reaches. It applies to any and every game in the cosmos....."
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30 of 36 people found the following review helpful
on March 28, 2011
Format: Hardcover
Martin Nowak is, first and foremost, a virtuoso. He has turned modelling into an art. Whether they deal with mutations inducing cancer, irregular verbs turning regular, the spread of HIV or the chemical lego of pre-life, his models always capture quintessential aspects with superb clarity and elegance. Even more remarkable as his light and confident touch in setting up the models is his dexterity in interpreting the results. He goes straight for the thrill. Nowak is enormously convincing and persuasive. This requires, of course, that he is convinced of himself. The evolution of cooperation has been the main theme of his stellar career, which led him in the briefest time from beginnings in Vienna to professorships in Oxford, Princeton and Harvard. Nowak's Hirsch-index is eighty (meaning that eighty of his more than three hundred papers have each been quoted in at least eighty papers), a fantastic achievement for someone in the mid-forties. Now he has decided to reach out for a larger audience, and written, with the expert support of Roger Highfield, one of Britain's foremost science writers, a book intended for the bestseller lists. Cooperation is a topic of central importance in many fields ranging from chemistry and biology to social and economic sciences, and Nowak is uniquely qualified, by his interdisciplinary background and his skills as communicator, to cover the whole canvas in a masterly tour.
One need not agree with all Nowak claims. In fact, some of the chapters will be highly controversial. Nowak defends his views with bravado, be they on kin selection or on Gustav Mahler's music. Some of these views make me slightly wince, but that is part of the fun. Nowak lives for his work, and merges with gusto his biography with the story of his field. Sobriety and a sceptical distance are for critics, not artists. Nowak runs on sheer enthusiasm, and conveys much of it to the reader. The book is a pleasure, heady, stimulating and brimful with adrenaline.
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5 of 5 people found the following review helpful
on January 27, 2013
Format: Paperback
The chatty writing, walks and hikes in woods and mountains, a ski hut, frequent major digressions, personal descriptions of grad student life, informal portraits of familiar names, interesting anecdotes, and indirect references to females, seemed eerily familiar. At first I wondered if I'd read this before. Then I realized, I'm reading the reincarnation of James D. Watson's "The Double Helix: A Personal Account of the Discovery of the Structure of DNA"! Except in that earlier book there had been only the one central bit of science, that science was already very famous and thoroughly described outside the book itself, and the author was quite young.

This book too is a personal description of scientific research. But here the description stretches over decades in different countries, the science itself --not all of it familiar-- is described in the same book, and there's a strong attempt to tie all the bits of science together under a single thematic umbrella. As a result the book unfortunately invites being addressed as something different than it is. Also, there's a constant unresolved tension between chronological and topical organizations.

The book can seem to be an overview of cooperation genetics, but mis-reading it this way is misleading. Significant contributions by the many other researchers in the field of course aren't covered at all. Some of the important first five chapters seemed on initial reading to be purely third-hand and awfully short (although to be fair, on checking back they turn out to be anchored by non-trivial research forays after all). Some research forays fit into the theme of cooperation only incidentally (ex: cooperation within groups requires communication between individuals, and the principal way humans communicate is with language, therefore this research on the origin of verbs is about cooperation -- ???). And there are gaps wherever Nowak himself hasn't delved into a sub-area. In the particular sub-area of enforcing cooperation through punishment, Nowak's interpretations are perhaps overstated, somewhat outside the mainstream, and frankly invite controversy.

I was initially drawn to this book by Nowak's reputation as a mathematician. I expected detailed mathematical descriptions of approaches to some of the thornier problems of cooperation genetics. I was sorely disappointed. There was apparently a confluence of some editor's "no equations" mantra with the author's dislike of complication. As a result not only is there no mathematics at all, but even the prose descriptions include what seemed to me an awful lot of hyperbole and impreciseness and even hand-waving.

High marks for the quite engaging writing style, the more than adequate organization, and the very strong and persistent effort to tie everything together into the cooperation theme. In these respects this book stands above most. There are hints sprinkled throughout the book that Nowak's research forays into cooperation genetics have been driven by preexisting personal philosophy, rather than the other way around. Although he scrupulously avoids any actual scientific whoppers, the overall feeling I got was of someone pushing a philosophical and religious agenda, or even of the dreaded "teleology". Because he's already shot this wad, the last chapter where I'd expect to find it is instead an unenlightening catalog of everything that ails us today. It felt more like liberal cocktail party chatter than the last chapter of a popular science book.
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14 of 18 people found the following review helpful
on February 14, 2012
Format: Kindle EditionVerified Purchase
I picked-up this book following a rather positive and good book review in the German version of the financial times. I will spare you the details of the topic as these are nicely described by other readers (and I've not finished reading the book yet). But essentially it describes how cooperating is a good thing (particularly if everybody in a society participates) and how things can go wrong if there just a few bad apples (defectors, cheats) that take advantage of everybody elses generosity. Good stuff.

What makes the book absolutely unreadable is the arrogance and self-centerdness of the author. First off, there are two authors yet there is always just the word "I"...."I came up with this...", "I published in Science Journal..." and in so many words this guy things he is the the greatest scientist the world has ever seen. Please bow to him now! It is incredibly irritating.

I write this as being a person in science and the one thing you realize is that the more you know the more humble you become because the world is an incredibly complex place. Math is great (I use it everyday), but honestly it becomes really painful to read over and over again how the authors mathematical models are able to predict everything...they can't, they don't, they never will. Whenever people say how great they are, particularly in science, you immediately become suspicious. Because truly outstanding people NEVER self-glorify. Never. Because if you do, you are not great. This "guy" is clearly not great.

His ego is soooo large than I'm surprised he has been allowed to even be at the various Institutions he constantly keeps mentioning. If there ever was a person that likes to name drop, he is it. Maybe he should move to the tabloid world?

So all in all, a great topic but an irritatingly arrogant author--he does not deserve your money. Read-up on the topic somewhere else.
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30 of 42 people found the following review helpful
on April 12, 2011
Format: Hardcover
Martin Nowak is a mathematician and scientist of great stature, expansive ambition, and all-embracing energy. He avoids being hog-tied to any particular discipline, but rather follows his fancy where it takes him in the behavioral sciences. Nowak is especially admirable because he is an iconoclast who comes out guns a-blazing to confront any Received Wisdom with which he fundamentally (or even marginally) disagrees. He apparently does not mind being wrong (he sometimes is) because when he is right, it more than makes up for his goof-ups. Nowak was a student of Peter Schuster, the great mathematical chemist, and a student and coauthor of the biological mathematician Karl Sigmund, at the University of Vienna in the late 1980's.

Nowak is currently the head of the Program for Evolutionary Dynamics at Harvard University. He was hired by Harvard after an admirer offer to contribute a Very Large Sum to the Harvard endowment as part of a deal to install him as Professor of Mathematics and Biology at the august institution. Nowak steps on lots of peoples toes, and there is little chance he would have been hired at the venerable school without the huge monetary emolument. This is no criticism of Nowak---Harvard does not cotton up to anti-Establishment, transdisciplinary, iconoclastic intellectuals. To round out Nowak's multidimensional cerebral life, he is an observant and pious Catholic, and his work at Harvard is co-sponsored by the Templeton Foundation, famous for its support of projects that link science and spirituality.

The main theme of SuperCooperators is both important and ubiquitous in modern behavioral science: we humans are who we are because we evolved that way biologically, and cooperation has been the key to our success as a species. "Out breathtaking ability to cooperate is one of the main reasons we have managed to survive in every ecosystem on Earth, from scorch, sun-baked deserts to the frozen wastes of Antarctica to the dark, crushing ocean depths" (the hyperbole is a typical of Nowak's literary tricks; in fact we have not survived in any of the three ecosystems he mentions, but we have in most of those he does not). If the last half of the Twentieth century was the Homage to Selfishness, the first half of the current century is the Homage to Cooperation. Perhaps the most important contribution to this new theme is John Maynard Smith and Eors Szathmary's The Origins of Life, which depicts each stage in the development of biological complexity as the synergistic synthesis of hitherto competing biological entities, from single-cell to multicellular organisms, from prokaryotic to eukaryotic cells, all the way up to social species, of which our species is among the most successful. As for humans, Samuel Bowles and I have argued this position for some fifteen years, our book "A Cooperative Species" (Princeton, 2011) coming out in a few months, and our colleagues Robert Boyd and Peter Richerson predated our effort by at least a decade.

SuperCooperators is extremely accessible to non-scientists. It dispenses with footnotes and highly accurate but complex phraseology, in favor of explaining concepts in a witty, conversational, and light-handed manner. One is struck throughout with Nowak's love of mathematics and science. He repeatedly conveys to the reader his the love of knowledge for its own sake (he regularly quotes from Albert Einstein, a man whose passion for truth approached theological levels). He also conveys his commitment to and fondness for his co-researchers, both teachers and mentors (the book is dedicated to Bob and Karl---presumably his mentor Robert May and his teacher and coauthor Karl Sigmund). In fact, SuperCooperators is much more a book about the work of Nowak and his Harvard Program for Evolutionary Dynamics than it is an even-handed overview of the field and its development. This was a good choice on Nowak's part, because it permits him to convey the degree of collegiality and personal commitment that is one of the great comforts of scientific research, and is often ignored in favor of maintaining the emotional flatness that is (falsely) reputed to be the hallmark of good science.

How do we account for human cooperation? Nowak offers five very broad mechanisms: direct reciprocity, indirect reciprocity, spatial selection, multilevel selection, and kin-selection. He devotes a chapter to each of these mechanisms, in each case highlighting his own contributions. This may sound self-aggrandizing, and it is. But I do not think this is a mistake. The benefit for readers is a sense of intimacy with the writer, who becomes the protagonist in a scientific melodrama, a knight in shining armor slaying falsity and promoting truth wherever his wanderings take him. However, this approach does not allow the reader to appreciate the substantive scientific issues involved. For instance, Nowak is strong supporter of kin-selection, but a bitter critic, along with Harvard colleague Edward O. Wilson and co-worker Corita Tarnita, of William Hamilton's explanation of kin-selection in terms of "inclusive fitness." Now you must understand, dear reader, that the inclusive fitness concept has become the standard explanation of cooperation in population biology and animal behavior theory, so Nowak's critique, presented in Nature in 2010, produced a major shake-up in the profession. Nowak presents his critique here, but does not attempt to relate the counterarguments, which are at least extremely interesting, and probably indicate serious problems with the Nowak-Tarnita-Wilson argument. For one think, the authors pose inclusive fitness theory as an "alternative to natural selection," which is pretty ludicrous. Right or wrong, inclusive fitness theory is a form of natural selection, not an alternative to natural selection.

To convey his light-handed treatment of scientific themes, consider Nowak's treatment of multilevel selection. After two short paragraphs going over the history of group selection (focus on Wynne-Edwards' latter years studying red grouse with binoculars from the window of a room in a rest home in the Dee Valley), Nowak confesses that earlier attempts to model multilevel selection were "too complicated for my taste" and that he and Karl Sigmund did not find them "particularly convincing." This manner of presenting objections to a scientific theory bypasses all substantive issues (what were the earlier models and what were their successes and deficiencies) and moves directly to the subjective emotions of Nowak and Sigmund. How does Nowak describe his and Sigmund's passage into the theory of multilevel selection. "In the heart of the Rauriser Urwald," he writes, "Karl [Sigmund] and I came across a little wooden board bearing a poem by Goethe (1749-1832)...Goethe's poem begins as follows: "Muesset beim Naturbetrachten immer eins wie alles acten." This translates as "When looking at nature, you must always consider the detail and the whole." I could not think of a better way to express the idea of multilevel selection and, though I realized the field had a long, troubled, and vexatious history, I began to think about whether it might work in the real world." Now, of course, the Goethe quote can be interpreted in many ways, but none of them has anything to do with multilevel or any other form of selection. Moreover, I frankly do not believe that this story is accurate. It sounds like an apocryphal yarn, like the one about Newton and the apple.

Nowak goes on to explain the model of multilevel selection with Arne Traulsen that is no different than a dozen previous models of the within-group cost and between-group benefit of altruistic behavior, and with no mention of Price's equation, which was the inspiration for all previous mathematical models of multilevel selection (Nowak belittles Price's equation later in the book as being "tautological", which is incorrect, as it depends on a empirically meaningful analytical representation of biological fitness). The paper Traulsen and Nowak wrote (PNAS 2006) was indeed simple and elegant, but hardly earthshaking. The elegance derives from its simplifying assumptions (recall Nowak's distaste of complexity), which include (a) no migration; (b) fixed maximum group size; (c) groups of maximum size consist of all cooperators or all defectors. None of these conditions is biologically relevant, so the Nowak Traulsen contribution is likely only of passing interest. To reiterate, I do not consider Nowak's self-aggrandizement a mistake, given his goal of presenting the non-scientist reader with a highly human treatment of behavioral and mathematical theory. The historians will sort out who actually did what, and the average reader does not really care at all. However, this approach does lead to the suppression of scientific content in favor of personal feelings and memorable anecdotes.

Given his strong personal commitments to his co-workers, it is not surprising that Nowak rarely has praise for the sustained and seminal contributions of contemporary researchers who work outside his research group. Luca Cavalli-Sforza, Robert Boyd, Peter Richerson, Robin Dunbar, Alan Grafen, Amos Zahavi, and Joseph Henrich are not mentioned at all, and others, including Samuel Bowles and Marcus Feldman, are mentioned in passing for a particular study or two, but nothing revealing the depth of their thought or nature of their contributions to modern behavioral science. Similarly, he has little room for contributions from other fields, such as economics, sociology, and psychology, or evidence from archeology, history, anthropology, or paleontology. This I also consider a shortcoming. Weaving the contributions of others with his own in a more substantive way would give the reader a better appreciation for the contemporary state of research on cooperation and conflict in the animal and human world. Perhaps this is just tit-for-tat, as I find that few authors from the behavioral sciences outside of Nowak's circle pay much attention to the work of Nowak and his students. My problem with Nowak's models is that they tend to remove important aspects of the subject being modeled in the interest of obtaining elegant formulas. What, for instance, is the value of a model of group selection without migration and variable group size? What, indeed, are we supposed to make of the Prisoner's Dilemma played on a grid, when in real life no species plays any games at all on a grid.

I like the fact that Nowak does not shy from headlong attack on ideas that he thinks are wrong. I am not so happy, however, when the ideas of people with whom I work are being attacked. Nowak's Chapter 12, "Punish and Perish," is just such an attack. In various works, my colleagues and I have proposed that humans exhibit a behavioral syndrome we call "strong reciprocity," according to which most humans have a predisposition to cooperate in collective endeavors that depend on voluntary participation, the success of which depends on a high rate of cooperation and a low rate of defection, and also have a predisposition to punish those who free-ride on the cooperation of others, without requiring that their efforts be repaid in the future. The latter predisposition has been called "altruistic punishment," and it is this concept to which Nowak objects. I think Nowak has correct objections to assertions we do not make but that he attributes to us, and incorrect objection concerning statements we do make.

Nowak begins Chapter 12 by saying that "Punishment is not, as some have claimed, a mechanism for the evolution of cooperation." Rather, he claims, "punishment fits neatly into the framework of the Prisoner's Dilemma." Here Nowak's claim is incorrect. The Prisoner's Dilemma, and direct reciprocity as well, are only relevant in dyadic interactions, and it is quite true that altruistic punishment is irrelevant in such interactions, or in fact in even broader interactions where reputation effects are powerful and ubiquitous. However, human cooperation occurs in large groups in which reputation effects have little or no power. It is these situations that altruistic becomes relevant for sustaining cooperation.

For instance, consider voting in a democratic society. It is costly to vote and because one vote cannot change the outcome of an election except in the smallest of communities, a purely self-regarding person would never vote. Voting is thus mostly an example of the positive side of strong reciprocity, that of altruistic cooperation. However, many people vote because their family, friends, and associates would disapprove of their lack of community spirit if they did not vote. If A does not vote and B expresses disapproval, then B incurs the cost of disrupting the normal relationship between A and B. Disapproving of A is thus an act of costly altruistic punishment, supporting democratic norms at personal expense without the likelihood of any personal gain therefrom. Without such altruistic punishment, voting might be considerably less common than it is, a situation that could delegitimize political democracy.

Consider also ostracism, which is a common group punishment to a member who violates the norms associated with group membership. An individual is ostracized when all (or at least most) group members rupture normal relations with the individual and/or actively prevent the individual from participating in group activities or sharing the benefits of these activities. We tend to ignore the altruistic punishment side of the ostracism phenomenon, but it is always present, unless there is an external third party who has an incentive to enforce the ostracism decision.

Nowak does recognize the existence of mufti-player games, as when he says "If I punish you for defecting in games with other players, that is indirect reciprocity." This statement, however is incorrect. Indirect reciprocity depends on the ability of the group to establish strong reputational information, but such information is generally not available in large groups. If I express to you my disapproval of your not voting who, other than yourself, will see this action and use it to increase my stock of "good reputation"? The answer is generally, no one; nor do I express my disapproval because I hope to gain reputation-wise.

Of course altruistic punishment is not a "mechanism of evolution" in the sense of direct reciprocity, indirect reciprocity, and the others among Nowak's "big five." The big five are social mechanisms, whereas, altruistic punishment is an evolved form of behavior exhibited by humans that contributes to human cooperation, and hence to the emergence of our species as a world-scale key player. It operates in direct interactions, large groups, and explains important aspects of multilevel selection in humans, but like big teeth, wings, or an advanced immune system, it is not a "mechanism" of evolution.

Nowak also stresses that in the many experiments that have shown the efficacy of altruistic punishment in fostering cooperation, in fact the cost of punishing often completely offsets the gains from cooperation. He does not, however, present the important study by GÃÃ,Ã,¤chter, Renner and Sefton (2008) in which subjects are allowed to interact over 50 periods rather than just 10. They found that after the initial rounds, the net benefits to the group with the punishment option significantly exceeded those of the no-punishment group with the difference in net payoffs growing over time, except for the final round in which the hapless end-game free-riders were heavily punished.

Given that most social dilemma interactions in neighborhoods, work teams, and the like, extend over far more than 10 periods, the concern that altruistic punishment lowers group benefits to be misplaced. The experiment to which Nowak refers (Dreber et al. 2008) does not constitute evidence for the counterproductive punishment hypothesis for the additional reason that their two-person game made punishment irrelevant, for one could always retaliate on a defector simply by withdrawing cooperation, thus obviating the need for any special kind of punishment. But while the 50-period design of the GÃÃ,Ã,¤chter et al. experiment corrects one of the design biases that suggested counterproductive punishment in the earlier experiment, their design still misses something essential to altruistic punishment in the real world: it is effective only if it is regarded as legitimate according to widely held social norms.

Ertan, Page, and Putterman (2009) designed an ingenious experiment to explore this possibility. They allowed experimental subjects prior to playing the public goods game to vote on whether punishment should be allowed and if so, should it be restricted in any manner. From their first opportunity to vote, no group ever allowed punishment of high contributors, most groups eventually voted to allow punishment of low contributors in the baseline treatments, and the result was both high contributions and high efficiency levels. In the laboratory, groups solved their free-rider problems by allowing low contributors alone to be punished. Apparently the determination of the punishment system by majority rule made the punishment not only an incentive but also a signal of group norms.

Nowak also argues that what we call altruistic punishment is often costly but not motivated at all by prosocial feelings of punishing norm violators. Rather it is motivated by spite and a taste for revenge. Punishing those who have hurt you is normally motivated by a taste for retribution rather than a selfless expression of support for public morality. Indeed, in our research we tend to term this costly punishment, and it is altruistic in the sense of being other-regarding. However, fully understand that retribution is generally not motivated by prosocial intentions. Much of the punishment that occurs under the rubric of strong reciprocity is motivated by the urge to retaliate simply because hurting those who hurt you is a selfish, however socially useful, pleasure.

Nowak's critique of altruistic punishment in Dreber et al. probably should not have been published because it is well known that cooperation can be sustained in the repeated prisoner's dilemma by self-regarding players without punishment (it is called tit-for-tat), and interpreting cooperating as "reward" in this setting is just a semantic ploy with no substance. However in 2009, in Science, Nowak published a true public goods game with punishment (groups of size four), called "Positive Interactions Promote Public Cooperation," with coauthors David G. Rand, Anna Dreber, Tore Ellingsen, and Drew Fudenberg. The paper contends that experimental evidence with human subjects in the laboratory shows that reward is more effective than punishment in eliciting cooperation in a public goods game.

The paper has two serious and obvious flaws. First it is indefinitely repeated, so there are no end game results. This design contrasts with virtually all previous studies of the public goods game, which assumes that players know exactly how many periods will be played, and much can be learned from the course of play from early to late rounds. It is well known, for instance, that the rate of decay of cooperation depends on the length of the game, and even when cooperation is sustained for most of the game, it can break down towards the final periods. In the Supplementary Online Material (SOM) the authors justify this deviation from the standard protocol by saying that indefinite repetition is more realistic than a fixed number of periods. This is true, but irrelevant. Having a fixed number of periods is an experimental control condition that is introduced precisely to avoid the ambiguity of indefinite repetition: there are an infinite number of equilibria to the indefinitely repeated game, and which if any is chosen is likely to depend on extraneous framing effects.

Much more important however, the "reward" in the experiment costs 4 points for the rewarding subject while the recipient gains 12 points. Of course reward will outperform punishment if the experimenter pays sufficiently for the reward, while the players pay for the punishment! It would be a surprising result if the authors came to the same conclusion with the recipient getting 4 points, not 12.

In fact, it is clear that the punishment/reward stages of the game in this paper form another public goods game with a multiplier of 3, in which individuals cooperate with no end-game effects. The fact that rewarding players can choose to whom to direct the benefits they generate is interesting, but we still have two overlapping public goods game in which the second includes a signal (previous contribution) that partially recreates the conditions of a two-player repeated prisoner's dilemma. This is a worthy subject of study, but in no way supports the authors' conclusions.

By the way, the reader would never suspect the existence of any of the egregious flaws in this paper by reading the published material. It is all available only in the Supplementary Online Material, which is read only by experts. I doubt that the reviewers of the paper even glanced at the Supplementary Online Material. If I were a Science editor, I would compel reviewers to review, separately, the main contribution and the supplementary material. The quality of reviewing might improve considerably, at least in areas with which I am acquainted.

Nowak's hostility to altruistic punishment appears to flow from his preference for rewarding good behavior rather than punishing bad behavior. This is of course very enlightened thinking when it comes to raising children and managing employees. But here he confuses the sorts of social dilemmas dealt with in the human cooperation literature with reward an punishment in the sort of principal-agent models that economists commonly deal with, such as employer employee. In an employer-employee setting, despite the predictions of standard economic theory, punishing bad behavior often has severely efficiency-reducing effects, whereas trusting an employee to "do the right thing" is often the most cost effecting strategy an employer can use. In effect, by trusting, the employer sets up a setting where the strong reciprocity predispositions of employees operate to great effect. This was shown by our colleagues Ernst Fehr. Simon Gaechter, and Georg Kirchsteiger (1997) in one of the most famous experiments in behavioral game theory, in which they show that trust is a potent contract enforcement device. More recently, Ernst Fehr and Bettina Rockenbach, in their paper "Detrimental Effects of Sanctions on Human Altruism", Nature 422 [March 13] (2003):137-140, showed clearly that when employers have an available punishment device but choose not to use it, they elicit the best performance from workers. Sanctions are important, but they surely backfire when deployed in inappropriate circumstances.

Social dilemmas where cooperation is voluntary and regulated by peer-relationships are not principal-agent models, which are dyadic hierarchical interactions. However, it is usually true even in principal-agent models that the principal has some means of punishing miscreant agents (e.g. firing them). Nowak's idea of a world in which reward reigns supreme and the threat of punishment is absent is not our world.

Nowak's iconoclasm is a wonderful gift to scientists, because he stirs up the muddy waters in fruitful directions even when he is ultimately wrong in his critiques. In the long run, however, Nowak's modus operandi may be costly to an assessment of his scientific contributions. Nowak often jumps into a field he barely knows, makes arguments that experts merely dismiss as half-baked, and is taken seriously only by scientists and non-scientists who are even more ignorant of the field than Nowak. Human cooperation is a case in point.
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8 of 11 people found the following review helpful
Format: Hardcover
This was an interesting book attempting to make accessible the basic ideas of game theory as they apply to evolution. In the end, though, I really think this book is badly organized, both within and between chapters.

First, the good: the first half of the book has some really interesting discussion about the key mechanism that allow (on the whole) selfish humans to engage in the act of cooperation. Direct reciprocity, indirect reciprocity, spatial games, group selection, and kin selection. Particularly illuminating are the discussions on why group selection has wrongly been dismissed by many as a valid evolutionary force (why, in other words, evolution can actually select groups in addition to individuals within groups), and why kin selection has wrongly been elevated as being more significant than it is (it is only significant if we assume that the relatives we are giving up our interests for actually WILL survive, rather than be subject to natural selection themselves). Also of interest was the section explaining why, contrary to what many think, tit for tat is not the best general strategy for virtually any variant of a prisoner's dilemma game. The second half of the book was also quite intriguing, looking at the evolution of cooperation from RNA sequences (pre-life, we're talking) to ant hives to humans' drive to punish defectors (at expense to themselves).

Now, the bad: the book was really disjointed. First, I was one - like a few other reviewers - who did not care for the author's frequent digression into details of his personal life. I, for one, don't really care that he thought of this idea while talking with x in a castle that had some really interesting objects in it. It doesn't add anything to the already interesting subject matter, and actually detracts from what could have been a nice flow. Also, while the two 'halves' of the book - one on the mechanisms we use for cooperation and the other on the origin of cooperation - are interesting, the two halves never really tie together. So, while the book's theme is indeed how self-interest has led humans (and some other species) to cooperate, the two halves really feel like two separate mini-books that never really tie together.

Overall, this book is pretty attention-holding, especially for those of us intrigued by how seemingly self-interested humans actually evolved as pretty good cooperators. (A book I might recommend after this one is Ridley's The Rational Optimist: How Prosperity Evolves (P.S.) or Wright's Nonzero: The Logic of Human Destiny, both of which are about how human societies have progressed the more they've discovered ways to engage in non-zero-sum transactions - cooperation.) But this book could have been organized and written better, using less personal digression and a bit more overall organization.

(Another really good book explaining game theory in a bit more depth (including the various games and how they can be seen in everyday life) to the general reader is Rock, Paper, Scissors: Game Theory in Everyday Life. I'd recommend that one before this one, even though they are on slightly different topics.)
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2 of 2 people found the following review helpful
on July 24, 2012
Format: HardcoverVerified Purchase
This is an excellent book,using Game Theory to argue how cooperation helped in the evolutionary process. I would have enjoyed it much better if not for the many unnecessary tales of how he met so-and-so, and the many anecdotal tales of the various scientists. Just stick to the science, and I would have given it a 5 stars.

I like particularly the last chapter where he summarized the whole book.

Well worth a read.
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2 of 2 people found the following review helpful
on April 13, 2013
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As a former research biologist, I was thrilled to find mathematical proof for the evolutionary value of cooperation under certain conditions. I was inspired to find and purchase Martin Nowak's previously published book, "Evolutionary Dynamics", which contains the math he used to write "Supercooperators".
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1 of 1 people found the following review helpful
on February 12, 2015
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Nowak is a respected Harvard professor of mathematical biology with numerous well regarded publications. Sadly, he has chosen to stray off the path into an arrogant attack on science motivated by religious fervor. His recent actions show the evil consequences when universities accept money from religious groups, science journals are so awed by big names that they avoid proper peer review, and egos are permitted to get out of control. Most of this book is good, but it repeats the utterly misconceived attacks on inclusive fitness in favor of group selection which he and his much more famous Harvard colleague E.O.Wilson have been making the last 5 years, so it is on that chapter of the book that I will concentrate. This takes us into the nature of evolution, the basics of scientific methodology, how math relates to science, what constitutes a theory, and what attitudes to religion and generosity are appropriate as we inexorably approach the collapse of industrial civilization.
Their attack on evolution came into general awareness in 2010 with the publication of a now infamous paper in nature which was almost universally rejected by over 140 biologists who responded with letters and many papers since have shown in detail why it is mistaken. Wilson and Nowak have not responded in any meaningful way in 5 years of articles, lectures and several books. There is no choice but to agree with Dawkin's trenchant comment "For Wilson not to acknowledge that he speaks for himself against the great majority of his professional colleagues is--it pains me to say this of a lifelong hero --an act of wanton arrogance." In view of Nowak’s subsequent behavior one must include him as well. Wilson has also been one of my heros and I feel like one of the stunned people one sees on TV being interviewed after the nice man next door, who has been babysitting everyone's children for 30 years, is exposed as a serial killer.
Dawkins also points out (as he and others have done for many years) that inclusive fitness is entailed by (i.e., logically follows from) neo-Darwinism and cannot be rejected without rejecting evolution itself. If you want a serious book length account of social evolution (i.e, inclusive fitness) by an expert see Principles of Social Evolution by Andrew F.G. Bourke.
After reading this book and its reviews, I dug into some of the scientific articles which responded to Nowak and Wilson and to Van Veelen’s critiques of the Price equation upon which they heavily relied. The reviews noted that it has always been clear that the math of group or multilevel selection reduces to that of inclusive fitness (kin selection) and that it is not logically possible to select for behavior that does not benefit the genes that are unique to the actor and its immediate relatives. To put it bluntly, ‘altruistic’ behavior is always selfish in the end in the sense that it increases survival of the genes in the altruist. This to me is obvious from daily life and any scientists who claim otherwise have clearly lost their way. Yes, it does happen in the weirdness of modern life (i.e., so unlike the stone age society in which we evolved) that one sometimes sees a person give their life to protect a nonrelated person, but clearly they will not do it again and (provided its done before they replicate) any tendency to do it will not be inherited either. Even if they have already replicated they will on average leave behind fewer descendants than if they held back. This guarantees that any genetic tendency for ‘true altruism’- i.e., behavior that decreases ones genes in the population-- will be selected against and no more than this very basic logic is needed to grasp evolution by natural selection, kin selection and inclusive fitness—all the mathematical niceties serving only to quantitate things and to clarify strange living arrangements in some of our relatives (e.g., ants, termites and mole rats).
The major focus of the group selectionist’s (‘groupies’) attack is the famous Extended Price Equation that has been used to model inclusive fitness, published by Price about 40 years ago. The best papers debunking these attacks that I have found are those of Frank and Bourke and I will start with a few quotes from Frank ‘Natural selection. IV. The Price equation’ J . EVOL. BIOL. 25 (2012) 1002–1019.
‘The critics confuse the distinct roles of general abstract theory and concrete dynamical models for particular cases. The enduring power of the Price equation arises from the discovery of essential invariances in natural selection. For example, kin selection theory expresses biological problems in terms of relatedness coefficients. Relatedness measures the association between social partners. The proper measure of relatedness identifies distinct biological scenarios with the same (invariant) evolutionary outcome. Invariance relations provide the deepest insights of scientific thought…Essentially, all modern discussions of multilevel selection and group selection derive from Price (1972a), as developed by Hamilton (1975). Price and Hamilton noted that the Price equation can be expanded recursively to represent nested levels of analysis, for example individuals living in groups… All modern conceptual insights about group selection derive from Price’s recursive expansion of his abstract expression of selection… A criticism of these Price equation applications is a criticism of the central approach of evolutionary quantitative genetics. Such criticisms may be valid for certain applications, but they must be evaluated in the broader context of quantitative
genetics theory…[and in a quote from Price … ‘Gene frequency change is the basic event in biological evolution. The following equation…which gives frequency change under selection from one generation to the next for a single gene or for any linear function of any number of genes at any number of loci, holds for any sort of dominance or epistasis, for sexual or asexual reproduction, for random or nonrandom mating, for diploid, haploid or polyploid species, and even for imaginary species with more than two sexes’…]… Path (contextual) analysis
follows as a natural extension of the Price equation, in which one makes specific models of fitness expressed by
regression. It does not make sense to discuss the Price equation and path analysis as alternatives… Critiques of the Price equation rarely distinguish the costs and benefits of particular assumptions in relation to particular goals. I use van Veelen’s recent series of papers as a proxy for those critiques. That series repeats some of the common misunderstandings and adds some new ones.

Nowak recently repeated van Veelen’s critique as the basis for his commentary on the Price equation (van Veelen, 2005; Nowak et al., 2010; van Veelen et al., 2010; Nowak& Highfield, 2011; van Veelen, 2011; van Veelen et al., 2012… This quote from van Veelen et al. (2012) demonstrates an interesting approach to scholarship. They first cite Frank as stating that dynamic insufficiency is a drawback of the Price equation. They then disagree with that point of view and present as their own interpretation an argument that is nearly identical in concept and phrasing to my own statement in the very paper that they cited as the foundation for their disagreement… The recursive form of the full Price equation provides the foundation for all modern studies of group selection and multilevel analysis. The Price equation helped in discovering those various connections, although there are many other ways in which to derive the same relations… Kin selection theory derives much of its power by identifying an invariant informational quantity sufficient to unify a wide variety of seemingly disparate processes (Frank, 1998, Chapter 6). The interpretation of kin selection as an informational invariance has not been fully developed and remains an open problem. Invariances provide the foundation of scientific understanding: ‘It is only slightly overstating the case to say that physics is the study of symmetry’ (Anderson, 1972). Invariance and symmetry mean the same thing (Weyl, 1983). Feynman (1967) emphasized that invariance is The Character of Physical Law. The commonly observed patterns of probability can be unified by the study of invariance and its association with measurement (Frank & Smith, 2010, 2011). There has been little effort in biology to pursue similar understanding of invariance and measurement (Frank, 2011; Houle et al.,2011).’

I hope it is becoming clear why I chose the title I did for this article. To attack the Price equation and inclusive fitness is to attack not only quantitative genetics and evolution by natural selection but the universally used concepts of covariance, invariance and symmetry which are basic to science and to rationality. Furthermore, the clearly voiced religious motivation of Nowak invites us to consider to what extent such Christian virtues as true (permanently genetically self-diminishing) altruism and the brotherhood of man (woman, child, dog etc.) can be part of a rational program for survival in the near future. My take is that true altruism is a luxury for those who don’t mind being evolutionary dead ends and that even in it’s ‘make believe’ inclusive fitness version one will be hard pressed to find it when the wolf is at the door (the inescapable scenario for the 12 billion in the next century).

There is much more in this gem which goes into exquisite logical and mathematical detail (and likewise his many other papers-you can get all 7 in this series in one pdf) but this will give the flavor. Another amusing episode concerns tautology in math. Frank again:
‘Nowak & Highfield (2011) and van Veelen et al. (2012) believe their arguments demonstrate that the Price equation is true in the same trivial sense, and they call that trivial type of truth a mathematical tautology. Interestingly, magazines, online articles and the scientific literature have for several years been using the phrase mathematical tautology for the Price equation, although Nowak & Highfield (2011) and van Veelen et al. (2012) do not provide citations to previous literature. As far as I know, the first description of the Price equation as a mathematical tautology was in the study of Frank (1995).’ Wittgenstein exposes a nearly universal misunderstanding of the role of math in science. All math (and logic) is a tautology that has no meaning or use until it is connected to our life with words. Every equation is a tautology until numbers and words and the system of conventions we call evolutionary psychology are employed. Amazingly Lamm in his recent excellent article ‘A Gentle Introduction to The Price Equation’ (2011) notes this:
‘The Price equation deals with any selection process. Indeed, we can define selection using it. It says nothing in particular about biological or genetic evolution, and is not tied to any particular biological scenario. This gives it immense power, but also means that it is quite possible to apply it incorrectly to the real world. This leads us to the second and final observation. The Price equation is analytic [true by definition or tautologous]. It is not a synthetic proposition [an empirical issue as to its truth or falsity]. We derived it based on straightforward definitions, and universal mathematical principles. The equation simply provides a useful way of interpreting the meaning of the straightforward definitions we started from. This however is not the case once you put the equation into words, thereby interpreting the mathematical relationships. If you merely say: _I define 'selection' to be the covariance blah blah blah,_ you might be safe. If you say: _the covariance blah blah blah is selection,_ you are making a claim with empirical content. More fundamentally, the belief that the rules of probability theory and statistics, or any other mathematical manipulation, describe the actual world is synthetic.’

So it should be clear that there is no such thing as sidestepping the Price equation and that like any equation it has limitless applications if one only connects it to the world with suitable words.

As Andy Gardner put it in his article on Price (Current Biology 18#5 R198). Also see his ‘Adaptation and Inclusive Fitness’ Current Biology 23, R577–R584, July 8, 2013
‘Such ideas were rather confused until Price, and later Hamilton, showed that the Price equation can be expanded to encompass multiple levels of selection acting simultaneously (Box 2). This allows selection at the various levels to be explicitly defined and separated, and provides the formal basis of group selection theory. Importantly, it allows the quantification of these separate forces and yields precise predictions for when group-beneficial behavior will be favoured. It turns out that these predictions are always consistent with Hamilton’s rule, rb – c > 0. Furthermore, because kin selection and group selection theory are both based upon the same Price equation, it is easy to show that the two approaches are mathematically exactly equivalent, and are simply alternative ways of carving up the total selection operating upon the social character. Irrespective of the approach taken, individual organisms are expected to maximize their inclusive fitness — though this result follows more easily from a kin selection analysis, as it makes the key element of relatedness more explicit.’

Consequently, to have the ‘groupies’ attacking the Price is bizarre. And here is Bourke’s recent summary of inclusive fitness vs ‘groupism’:
(haplodiploid and eusocial refer to the social insects which provide some of the best tests).

‘Recent critiques have questioned the validity of the leading theory for explaining social evolution and eusociality, namely inclusive fitness (kin selection) theory. I review recent and past literature to argue that these critiques do not succeed. Inclusive fitness theory has added fundamental insights to natural selection theory. These are the realization that selection on a gene for social behaviour depends on its effects on co-bearers, the explanation of social behaviours as unalike as altruism and selfishness using the same underlying parameters, and the explanation of within-group conflict in terms of non-coinciding inclusive fitness optima. A proposed alternative theory for eusocial evolution assumes mistakenly that workers’ interests are subordinate to the queen’s, contains no new elements and fails to make novel predictions. The haplodiploidy hypothesis has yet to be rigorously tested and positive relatedness within diploid eusocial societies supports
inclusive fitness theory. The theory has made unique, falsifiable predictions that have been confirmed, and its evidence base is extensive and robust. Hence, inclusive fitness theory deserves to keep its position as the leading theory for social evolution.’

However inclusive fitness (especially via the Extended Price Equation) explains much more than ant society, it explains how multicellular organisms came into being.

‘The third insight of inclusive fitness theory is the demonstration that conflict between members of a society is potentially present if they are unequally related to group offspring, since differential relatedness leads to unequal inclusive fitness optima . From this has sprung an understanding of an immense range of kin-selected conflicts, including conflicts within families and eusocial societies and intragenomic conflicts that follow the same underlying logic. The corollary of this insight is that societies are stable to the extent that the inclusive fitness optima of their members coincide. This in turn provides the rationale for the entire ‘major transitions’ view of evolution, whereby the origin of novel types of group in the history of life (e.g. genomes within cells, multicellular organisms and eusocial societies) can be explained as the result of their previously independent constituent units achieving a coincidence of inclusive fitness optima through grouping . From this standpoint, a multicellular organism is a eusocial society of cells in which the members of the society happen to be physically stuck together; the more fundamental glue, however, is the clonal relatedness that (barring mutations) gives each somatic cell within the organism a common interest in
promoting the production of gametes…Nowak et al. argued that their perspective assumes a ‘gene-centred approach’ that ‘makes inclusive fitness theory unnecessary’. This is puzzling, because entirely lacking from their perspective is the idea, which underpins each of inclusive fitness theory’s insights, of the gene as a self-promoting strategist whose evolutionary interests are conditional on the kin class in which it resides…In their model of the evolution of eusociality, Nowak et al. deduced that the problem of altruism is illusory. They wrote that ‘There is no paradoxical altruism that needs to be explained’ because they assumed that potential workers (daughters of a colony-founding female or queen) are ‘not independent agents’ but rather can be seen ‘as “robots” that are built by the queen’ or the ‘extrasomatic projection of [the queen’s] personal genome’. If this claim were correct, then only the queen’s interests would need to be addressed and one could conclude that worker altruism is more apparent than real. But it is incorrect, for two reasons. One is that, as has repeatedly been argued in response to previous ‘parental manipulation’ theories of the origin of eusociality , the inclusive fitness interests of workers and the mother queen do not coincide, because the two parties are differentially related to group offspring . The second is that worker behaviours such as eating of the queen’s eggs, egg-laying in response to perceived declines in queen fecundity, sex-ratio manipulation by destruction of the queen’s offspring and lethal aggression towards the queen all demonstrate that workers can act in their own interests and against those of the queen. In the light of this proven lack of worker passivity, workers’ reproductive self-sacrifice is paradoxical at first sight and this is the genuine problem of altruism that inclusive fitness theory has solved. (c) Alternative theory of eusocial evolution. Nowak et al. [38] presented an ‘alternative theory of eusocial evolution’ (as alluded to in §2b), backed up by a ‘mathematical model for the origin of eusociality’. However, these do not represent true alternative theories, either alone or in combination, because they do not make any points or predictions that have not been made within inclusive fitness theory’

Speaking of various steps in a scheme suggested by Nowak et al, Bourke says:
‘These steps constitute a reasonable scenario for the origin and elaboration of insect eusociality, but neither the sequence of steps nor the individual elements differ substantially from those that have been proposed to occur within the inclusive fitness framework…The alternative theory of eusocial evolution of Nowak et al. also exhibits two important weaknesses. To begin with, by allowing groups to form in multiple ways in step (i) (e.g. subsocially through parent–offspring associations but also by any other means, including ‘randomly by mutual local attraction’), their scenario ignores two critical points that are inconsistent with it but consistent with inclusive fitness theory. First, the evidence is that, in almost all eusocial lineages, eusociality has originated in social groups that were ancestrally subsocial and therefore characterized by high within-group relatedness. Second, the evidence is that the origin of obligate or complex eusociality, defined as involving adult workers irreversibly committed to a worker phenotype, is associated with ancestral lifetime parental monogamy and hence, again, with predictably high within-group relatedness…In sum, Nowak et al. make a case for considering the effect of the population-dynamic context in which eusocial evolution occurs. But their alternative theory and its associated model add no fundamentally new elements on top of those identified within the inclusive fitness framework and, relative to this framework, exhibit substantial shortcomings…More fundamentally, as has long been recognized and repeatedly stressed , the haplodiploidy hypothesis is not an essential component of inclusive fitness theory, since Hamilton’s rule for altruism can hold without the relatedness asymmetries caused by haplodiploidy being present. Highlighting the status of the haplodiploidy hypothesis to criticize inclusive fitness theory therefore misses the target. It also overlooks the fact that all diploid eusocial societies identified since the haplodiploidy hypothesis was proposed have turned out to be either clonal or family groups and so, as predicted by inclusive fitness theory, to exhibit positive relatedness. This is true of ambrosia beetle, social aphids, polyembryonic wasps, social shrimps and mole-rats. It is even true of a newly discovered eusocial flatworm . In short, the diploid eusocial societies, far from weakening inclusive fitness theory, serve to strengthen it…More broadly, the theory uniquely predicts the absence of altruism (involving lifetime costs to direct fitness) between non-relatives, and indeed no such cases have been found except in systems clearly derived from ancestral societies of relatives. Finally, inclusive fitness theory is unique in the range of social phenomena that it has successfully elucidated, including phenomena as superficially dissimilar as the origin of multicellularity and the origin of eusociality, or intragenomic conflicts and conflicts within eusocial societies. Overall, no other theory comes close to matching inclusive fitness theory’s record of successful explanation and prediction across such a range of phenomena within the field of social evolution. The challenge to any approach purporting to replace inclusive fitness theory is to explain the same phenomena without using the insights or concepts of the theory…Recent critiques of inclusive fitness theory have proved ineffective on multiple fronts. They do not demonstrate fatal or unrecognized difficulties with inclusive fitness theory. They do not provide a distinct replacement theory or offer a similarly unifying approach. They do not explain previously unexplained data or show that explanations from inclusive fitness theory are invalid. And they do not make new and unique predictions. The latest and most comprehensive critique of inclusive fitness theory, though broad-ranging in the scope of its criticism, suffers from the same faults. Certainly, relatedness does not explain all variation in social traits . In addition, the long-standing message from inclusive fitness theory is that particular combinations of non-genetic (e.g. ecological) and genetic factors are required for the origin of eusociality. Nonetheless, relatedness retains a unique status in the analysis of eusocial evolution because no amount of ecological benefit can bring about altruism if relatedness is zero.’
Andrew F. G. Bourke The validity and value of inclusive fitness theory Proc. R. Soc. B 2011 278, doi: 10.1098/rspb.2011.1465 14 September(2011)

One thing rarely mentioned by the groupies is the fact that, even were ‘group selection’ possible, selfishness is at least as likely (probably far more likely in most contexts) to be selected for as altruism. Just try to find examples of true altruism in nature –the fact that we can’t (which we know is not possible if we understand evolution) tells us that its apparent presence in humans is an artefact of modern life concealing the facts and that it can no more be selected for than the tendency to suicide (which in fact it is). One might also benefit from considering a phenomenon never (in my experience) mentioned by groupies--cancer. No group has as much in common as the (originally) genetically identical cells in our own bodies-a 100 trillion cell clone-- but we all born with thousands and perhaps millions of cells that have already taken the first step on the path to cancer and generate millions to billions of cancer cells in our life. If we did not die of other things first, we (and perhaps all multicellular organisms) would all die of cancer. Only a massive and hugely complex mechanism built into our genome that represses or derepresses trillions of genes in trillions of cells, and kills and creates billions of cells a second, keeps (the majority of us) alive long enough to reproduce. One might take this to imply that a just, democratic and enduring society for any kind of entity on any planet in any universe is only a dream, and that no being or power could make it otherwise. It is not only ‘the laws’ of physics that are universal and inescapable, or perhaps we should say that inclusive fitness is a law of physics.

In a bizarre twist, it was apparently such thoughts that drove Price (creator of the Price equation and a devout Christian) to suicide.

Many beside groupies note the pleasant nature of much human interaction and see a rosy future ahead-- but they are blind. It is crushingly obvious that the pleasantry is a transient phase due to abundant resources produced by the merciless rape of the planet, and as they are exhausted in the next two centuries or so, there will be misery and savagery worldwide as the (likely) permanent condition. Not just movie stars, politicians and the religious are oblivious to this, but even very bright academics who should know better. In his recent book ‘The Better Angels of Our Nature’ one of my most admired scholars Steven Pinker spends half the book showing how we have gotten more and more civilized, but he seems never to mention the obvious reasons why--the temporary abundance of resources coupled with massive police and military presence facilitated by surveillance and communication technologies. As industrial civilization collapses, it is inevitable that the Worst Devils of Our Nature will reappear. One sees it in the current chaos in the Middle East and Africa and even the world wars were Sunday picnics compared to what’s coming. Perhaps half of the 12 billion then alive will die of starvation, disease and violence, and it could be many more.

The facts that Wilson, Nowak et al have for four years persisted in publishing and making extravagant claims for grossly inadequate and thoroughly debunked work is not the worst of this scandal. It turns out that Nowak’s professorship at Harvard was purchased by the Templeton Foundation—well known for its pervasive sponsorship of lectures, conferences and publications attempting to reconcile religion and science. Nowak is a devout Catholic and it appears that a large gift to Harvard was contingent on Nowak’s appointment. Any of the groupies could have gone to the experts to learn the error of their ways (or just read their papers). But Nowak has received something like $14 million in Templeton grants in a few years and who wants to give that up?

Regarding Nowak and Highfield’s book ‘SuperCooperators’ I will let Dawkins do the honors:
‘I have read the book by Nowak and Highfield. Parts of it are quite good, but the quality abruptly, and embarrassingly, plummets in the chapter on kin selection, possibly under the influence of E O Wilson (who has been consistently misunderstanding kin selection ever since Sociobiology, mistakenly regarding it as a subset of group selection). Nowak misses the whole point of kin selection theory, which is that it is not something additional, not something over and-above ‘classical individual selection’ theory. Kin selection is not something EXTRA, not something to be resorted to only if ‘classical individual selection’ theory fails. Rather, it is an inevitable consequence of neo-Darwinism, which follows from it deductively. To talk about Darwinian selection MINUS kin selection is like talking about Euclidean geometry minus Pythagoras’ theorem. It is just that this logical consequence of neo-Darwinism was historically overlooked, which gave people a false impression that it was something additional and extra. Nowak’s otherwise good book is tragically marred by this elementary blunder. As a mathematician he really should have known better. It seems doubtful that he has ever read Hamilton’s classic papers on inclusive fitness, or he couldn’t have misunderstood the idea so comprehensively. The chapter on kin selection will discredit the book and stop it being taken seriously by those qualified to judge it, which is a pity.’ You can find this on the site whyevolutionistrue 2011/03/16.

A scathing review of ‘SuperCooperators’ also appeared from eminent game theorist/economist/political scientist (and Harvard alumnus) Herbert Gintis (who recounts the Templeton scandal therein), which is quite surprising considering his own love affair with group selection—see the review of his book with Bowles by Price – 2012. 10(1): 45-49.

In the end it is clear that this whole sad affair will be only the tiniest bump on the road and, like all things which exercise our attention now, will soon be forgotten as the horrors of unrestrained motherhood bring society crashing down. But one can be sure that even when global warming has put Harvard beneath the sea and starvation, disease and violence are the daily norm, there will be those who insist that it is not due to human activities (the opinion of half the American public currently) and that overpopulation is not a problem (the view of 40%), and there will be billions praying to their chosen deity for a rain of Big Macs from the sky, and that (assuming the enterprise of science has not collapsed, which is assuming a lot) someone somewhere will be writing a paper embracing group selection.
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